Blumea 55, 2010: 105–110 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE doi:10.3767/000651910X525259

A revision of Madagascan Bertiera (Rubiaceae)

P.M. Wittle1, A.P. Davis1

Key words Abstract A taxonomic revision of the Madagascan representatives of the genus Bertiera is presented, to include three species, B. brevithyrsa, B. crinita and B. longithyrsa. A key to their identification is provided, each species is Bertiera fully described, and summaries of distribution, habitat and ecology, and phenology are given; conservation assess- Bertiereae ments are also provided. Bertiera brevithyrsa is described as new species; B. crinita represents a new combination, conservation and a lectotype is designated for this name. IUCN Red List Categories Madagascar Published on 5 August 2010 Rubiaceae

INTRODUCTION MATERIALS AND METHODS

The genus Bertiera Aubl. comprises c. 52 species, and occurs This study is based on herbarium specimens and associated from Mexico to tropical America, tropical Africa, Madagascar spirit material, at the herbaria of K, MO, P, TAN and TEF (ab- and the Mascarenes, with the greatest species diversity in Africa breviations after Holmgren et al. 1990). Each specimen was (www.rbgkew.org.uk/data/rubiaceae). The tribal placement databased, and where possible georeferenced. Ecological of Bertiera has been the matter of recent systematic debate, and geographical data were collected from specimen labels. with placements in the tribe Heinsieae (Verdcourt 1983: 549), The measurements provided in the descriptions were made Gardenieae DC. subtribe Gardeniinae (Robbrecht et al. 1994), from herbarium specimens and spirit material. All material Gardenieae subtribe Bertierinae K.Schum. (Bridson 1998: was examined using a Leica MZ95 stereomicroscope. The 214), Coffeeae DC. (Andreasen & Bremer 2000), Bertiereae georeferenced specimen data was imported into ArcView™ (K.Schum.) Bridson (Bridson & Verdcourt 2003: 386) and Cof- geographic information software to produce distribution maps, feeae subtribe Bertierinae (K.Schum.) Robbr. & Manen (Rob- ecological data and to calculate area occupancy (AOO) and brecht & Manen 2006). Recent molecular and morphological extent of occurrence (EOO) for each species. AOO and EOO data (Davis et al. 2007, Tosh et al. 2009) supports placement figures were used in conjunction with field observations to within its own tribe, Bertiereae (Bridson & Verdcourt 2003). produce conservation ratings based on the IUCN Red List The only complete revision of Bertiera (Wernham 1912) is Categories criteria (IUCN 2001) using the methodology of now very much out of date, although useful works exist for Willis et al. (2003). Climatic data was extracted from BIOCLIM America (Steyermark 1967), Africa (Bridson 1988, Hallé 1960, data (Hijmans et al. 2005), and elevation data from the Digital 1970, Robbrecht et al. 1994, Nguembou et al. 2003) and the Elevation Model (DEM). Macarenes (Verdcourt 1983, Leroy 1989). For Madagascar there is presently only a single species known, B. longithyrsa REMARKS ON THE MORPHOLOGY OF BERTIERA IN Baker, which according to Baker (1890) closely resembles the MADAGASCAR Mauritian species B. zaluzania Gaertn. Following this work little has been written about B. longithyrsa, or Bertiera in Madagas- Bertiera brevithyrsa, B. crinita and B. longithyrsa sit firmly within car. Robbrecht et al. (1994) briefly mentions B. longithyrsa, and the circumscription of Bertiera (Wernham 1912, Robbrecht et commented that it can be separated from other Bertiera species al. 1994, Bridson & Verdcourt 2003), possessing the salient by its conspicuous sub-foliaceous bracts. Schatz (2001) treats characters of the genus: stipule pairs shortly connate above Madagascan Bertiera, comprising one species (B. longithyrsa), each node, inflorescences terminal (although rarely axillary; but infers that there are other species present in Madagascar Leroy 1974, Robbrecht et al. 1994); corolla lobes contorted (“2 or more? spp.”). The original description of B. longithyrsa to the left; stigma (pollen presenter) 10-winged/ridged in the remains the most comprehensive work on Madagascan Ber- upper part; ovary 2-locular; placentas peltate, with numer- tiera. Upon examination of material held at several herbaria ous ovules over the entire surface of each placenta; fruits (see Materials and Methods) and on the basis of fieldwork, it berry-like, containing numerous angular seeds (surface finely became apparent that a revision of the Madagascan species reticulate/pitted). Bertiereae comprises a single genus, and so was needed. In particular, it is necessary to recognize two the salient characters of Bertiera are the same as those of the other species of Bertiera for Madagascar, including: B. crinita tribe (Bridson & Verdcourt 2003, Davis et al. 2007). Overall, (A.Rich.) Wittle & A.P.Davis, a new combination based on the Madagascan species are similar to the Mascarene spe- Sabicea crinita A.Rich., and B. brevithyrsa A.P.Davis, a new cies (B. bistipulata Bojer, B. borbonica A.Rich. ex DC., B. rufa species from NE Madagascar. A.Rich. ex DC. and B. zaluzania Comm. ex C.F.Gaertn). In the three Madagascan species each flower is subtended (i.e. at the base of the hypanthium) by a pseudo-whorl of two to four 1 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, (rarely five) bracteoles, which are broader and more conspicu- UK; corresponding author e-mail: [email protected]. ous in B. crinita than B. brevithyrsa and B. longithyrsa. Flower

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Nothing in this license impairs or restricts the author’s moral rights. 106 Blumea – Volume 55 / 2, 2010 subtending bracteoles are present in the Mascarene species Phenology — Flowering: (August–) September to March; but they are fewer and smaller (Verdcourt 1983, Leroy, 1989), fruiting: November to May. or sometimes absent. KEY TO SPECIES TAXONOMIC TREATMENT FOR MADAGASCAN BERTIERA 1. Young shoots pubescent to densely pubescent (hairs 0.7–3 mm long); lower leaf surface (including secondary and terti- Bertiera Aubl. (1775a) 180; (1775b) t. 69; Wernham (1912) 110; N.Hallé ary venation) sparsely pubescent (hairs 0.4–1.7 mm long) (1960) 280. — Type species: Bertiera guianensis Aubl...... 2. B. crinita 1. Young shoots puberulous to pubescent (hairs 0.3–0.8 mm Description of Madagascan Bertiera long); lower leaf surface (including secondary and tertiary Shrubs or small trees. Many vegetative parts (shoots, stipules, venation) sparsely puberulous (hairs 0.2–0.3 mm long) 2 petioles, leaf margins, leaf midrib and other venation), puberu- 2. Leaf blades 9–22.5 by 2–7 cm; inflorescence an open, lous to densely pubescent. Young shoots terete. Stipules pairs rather lax thyrse, 4.5–20 cm long, peduncle 1–4.9 cm long; shortly connate (united in basal 1/3) above each node or ± free, bracteoles narrowly triangular to narrowly lanceolate, 0.5–1 narrowly triangular to ± lanceolate; apex long acuminate. mm wide. — NW Madagascar . . . . 3. B. longithyrsa Leaves petiolate; leaf blades narrowly elliptic to elliptic, elliptic- 2. Leaf blades 4–10.4 by 1.1–2.9 cm; inflorescence a compact oblong, or elliptic-obovate to elliptic-ovate, or narrowly obovate; thyrse, 1–2 cm long, peduncle 0.2–0.4 cm long; bracteoles base narrowly cuneate to cuneate-rounded; apex attenuate to linear to linear-triangular, 0.2–0.4 mm wide. — NE Mada- attenuate-acuminate, or sometimes narrowly acute, charta- gascar ...... 1. B. brevithyrsa ceous; margins flat; secondary venation eucamptodromus to weakly brochidodromus, 6–11 pairs; tertiary venation ramified to reticulate, indistinct to invisible; higher order venation reticulate 1. Bertiera brevithyrsa A.P.Davis, spec. nov. — Map 1 to ramified obscure to ± invisible; domatia present or absent, Bertierae longithyrsae Baker affinis sed foliis 4–10.4 longis 1.1–2.9 cm latis if present found in the axils of the midrib and secondary vein (nec 9–22.5 longis neque 2–7 cm latis), inflorescentia 1–2 cm (nec 4.5–20 on the lower surface of the leaf, pit-like, 0.2–0.5 mm diam, cm) longo, pedunculo 0.2–0.4 cm (nec 1–4.9 cm) longo, bracteolis linearis usually obscured by the hairs of the midrib and the second- vel lineari-triangularis, 0.2–0.4 mm (nec anguste triangularis neque anguste ary veins; on the upper surface usually not visible, if present lanceolatis, 0.5–1 mm) latis, differt. — Typus: Davis & Rakotonasolo APD 4524 (holo K; iso BR, MO, P, TAN, TEF), Madagascar, Province Toamasina, manifest as an obscure pustule-like swelling. Inflorescence a Region Analanjirofo, District Maroantsetra, Makira Protected Area, c. 15 km terminal thyrse, open (rather lax) or compact (inflorescence due west of Ambinanitelo, near Mt Beanivona, 16 May 2007. branches short), pedunculate, all parts (peduncle, inflorescence branches, bracts and bracteoles) puberulous to pubescent; Shrub or small narrow tree, 3–5 m high. Young shoots pu- bracts present or absent, in axils of first order inflorescence berulous to pubescent (hairs 0.3–0.5 mm long, light brown). branches, single, paired only if inflorescence branches oppo- Stipules 8–12.6 by 2.6–3.8 mm, pubescent (young stipules) site, linear, narrowly triangular, or very narrowly elliptic to very to sparsely pubescent or with hairs mainly confined to central narrowly oblanceolate; bracteoles subtending the flower (at the portion of stipule (hairs 0.4–0.7 mm long, brown) and the rest base of hypanthium), in groups (pseudo-whorls) of 2–4(–5), of the stipule glabrous to puberulous. Leaves: petioles 8–12.4 within groups size unequal, very narrowly triangular to ± lan- mm long, pubescent (like the young shoots); leaf blades nar- ceolate, or linear to linear-triangular. Flowers hermaphroditic, rowly obovate to narrowly elliptic, 4–10.4 by 1.1–2.9 cm; base homostylous, secondary pollen presentation present, sessile, narrowly cuneate, slightly attenuate, rarely slightly asymmetric; 5-merous, flower buds with corolla lobes overlapping to the margin puberulous to very sparsely pubescent (hairs 0.2–0.5 left; hypanthium ± obconical to urceolate, puberulous to pubes- mm long, light brown); apex attenuate to attenuate-acuminate; cent; calyx tube short, 0.5–1 mm long; calyx lobes triangular lower surface: midrib pubescent (hairs 0.4–0.5 mm long, ap- to narrowly triangular, or lanceolate, apices narrowly acute; pressed, light brown to brown); secondary veins (6–)7–8(–9) corolla ± tubular-infundibuliform, constricted in the upper 1/3 pairs, pubescent (like the midrib but density and length of hairs and expanded above the constriction, white (greenish white, less); tertiary venation ramified to reticulate, rather obscure; yellowish white, cream-white), slightly fleshy, external surface higher order venation obscure; leaf surface (including second- sparsely puberulous to sparsely pubescent, internal surface ary and tertiary venation) sparsely puberulous (hairs 0.2–0.3 densely pubescent in apical 1/3 (hairs white); corolla lobes mm long, brown); upper surface: midrib sparsely pubescent triangular to narrowly triangular; disc ± cylindrical, flat-topped, (hairs 0.4–0.6 mm long, light brown), venation and leaf surface glabrous; stamens: anthers included, fixed within the upper glabrous. Inflorescence a compact thyrse, 1–2 by 1–1.8 cm, all 1/4 of the corolla tube, introrse, sessile, ± medifixed, very nar- parts (peduncle, inflorescence branches, bracts and bracteoles) rowly ellipsoid-oblong, apex and base acute, with a brownish sparsely pubescent to pubescent (hairs appressed, 0.3–0.5 (when dry) apical connective appendage; ovary ovoid to ovoid- mm long, light brown to brown); peduncle 0.2–0.4 cm long; spherical, bilocular, placentation axile, placenta fixed ± at the bracts present or absent, if present narrowly triangular to linear, midpoint, distinctly peltate, with numerous ovules over entire 5.5–7.4 by 0.3–0.5 mm; bracteoles linear to linear-triangular, surface of each placenta; style narrowly club-shaped in general 5.1–7.2 by 0.2–0.4 mm. Flowers: hypanthium 1.4–1.7 by outline; stigma (pollen presenter) included, 10-winged/ridged, 1.1–1.4 mm, pubescent (hairs 0.3–0.5 mm long, whitish brown apex acute. Fruits berry-like, ellipsoid to ovoid, green to white to light brown); calyx lobes very narrowly triangular, 2.8–3.2 by (immature) to white and blue (nearly mature), then blue to dark 0.3–0.5 mm; other flower parts unknown. Fruit (4.2–)6.1–6.8 blue (mature), external surface glabrous to pubescent; calyx by 4.2–6.6 mm, sparsely pubescent (hairs 0.3–0.4 mm long, lobes persistent. Seeds flattened-angular, surface minutely whitish brown to light brown). Seeds 0.6–0.9 by 0.2–0.6 by reticulate/pitted, brown (when dry). 0.6–0.7 mm. Distribution — NW and E Madagascar (Map 1). Distribution — NE Madagascar. Province Toamasina, Re­ Habitat & Ecology — Occurring in humid, evergreen for- gion Analanjirofo (District Maroantsetra). est and seasonally dry evergreen-deciduous forest, mostly Habitat & Ecology — Based on specimen data: closed can­ in primary forest but also recorded in secondary vegetation; opy, humid evergreen forest, riverine vegetation, with only slight 0–1300(–1500) m. disturbance; recorded on quartz; c. 700 m. Based on GIS data: P.M. Wittle & A.P. Davis: A revision of Madagascan Bertiera 107 humid forest; igneous and metamorphic rocks; 900 m; mean 2. Bertiera crinita (A.Rich.) Wittle & A.P.Davis, comb. nov. annual precipitation 2 000 mm; mean annual temperature: — Fig. 1; Map 1 21 °C; dry season 0(–2) months. Basionym: Sabicea crinita A.Rich. (1830) 148. — Mussaenda crinita (A.Rich.) Phenology — Poorly known. Flowering: unknown; fruiting: Homolle (1938) 3. — Type: Herb. Richard s.n. [large specimen on sheet, May. excluding small specimen of a Mussaenda] (lectotype P, designated here), Vernacular names — None known. Madagascar, without date. Conservation status — IUCN Red List Category (IUCN 2001): Data Deficient (DD). Bertiera brevithyrsa is only known from two Shrub or small narrow tree, 1–5 m high. Young shoots pubes- collections, from a single locality/population. Further fieldwork cent to densely pubescent (hairs 0.7–3 mm long, ginger-brown). in the Makira Protected Area and nearby forested areas in Stipules 10.8–23.4 by 2.9–7.4 mm, sparsely to densely pubes- north eastern Madagascar is required before a conservation cent (hairs 1–3.1 mm long, ginger-brown). Leaves: petioles assessment can be made. (1–)3–10.6 mm long, pubescent (like the young shoots); leaf Note — This species was first discovered in 2007, from the blades narrowly elliptic to elliptic, elliptic-oblong, or elliptic- Makira Protected Area, near Maroantsetra in NE Madagascar. It ovate, 7.5–16.5 by 2–5.3 cm; base narrowly cuneate to ± is so far known only from a single locality within Makira, despite rounded, rarely slightly asymmetric; margin sparsely pubescent exploration in several localities in the same protected area. For (hairs 0.6–1.7 mm long, ginger-brown to brown); apex attenu- details of the difference between B. brevithyrsa and B. crinita ate to attenuate-acuminate; lower surface: midrib pubescent to see Notes for the latter species (below). See Key to Species densely pubescent (hairs 0.4–2.2 mm long, erect to appressed, for differences between B. brevithyrsa and B. longithyrsa. light brown to ginger-brown); secondary veins 7–11 pairs, in- tersecondaries sometimes present, pubescent (like the midrib Additional material. Briggs 125 (K, MO, P, TAN, TEF), Madagascar, but density and length of hairs less); tertiary venation ramified Province Toamasina, Region Analanjirofo, District Maroantsetra, Makira Pro- tected Area, c. 15 km due west of Ambinanitelo, near Mt Beanivona (Andra- to reticulate, rather obscure or ± absent; higher order venation menahely camp), 14 May 2007. obscure to ± invisible; leaf surface (including secondary and tertiary venation) sparsely pubescent (hairs 0.4–1.7 mm long, brown to ginger-brown); upper surface: midrib, venation and leaf surface sparsely pubescent (like the lower surface but density and length of hairs less to much less), venation and leaf surface very rarely glabrous. Inflorescence a compact thyrse, rarely rather open and lax but usually becoming open and lax at fruiting, 1.5–12.9(–16.8) by 1.8–6.3 cm, all parts (peduncle, inflorescence branches, bracts and bracteoles) pubescent to densely pubescent (hairs erect to erecto-patent, (0.4–)0.7–1.7 mm long, white-brown to brown, or ginger-brown); peduncle 0.2–3.5(–5.2) cm long; bracts present or absent, if present narrowly triangular, linear, or rarely very narrowly elliptic, 6.7– 13.4(–25.8) by 0.4–1.8(–3.2) mm; bracteoles very narrowly triangular to ± lanceolate, 3.9–8.9 by 0.5–1.5 mm. Flowers: hypanthium 2.8–3.5 by 1.4–1.6 mm, sparsely pubescent (hairs 0.3–0.7 mm long, brown to ginger-brown); calyx lobes trian- gular to narrowly triangular or lanceolate, 1–4 by 0.4–0.7 mm; corolla 6.1–12.5 by 1.8–3.7 mm, exterior sparsely pubescent, hairs mainly confined to corolla lobes (hairs c. 0.5 mm long, light brown); corolla tube 2.5–8 by 1.6–2.5 mm; corolla lobes narrowly triangular, 1.9–4.7 by 0.7–1.7 mm, apices acute to acuminate, acumen c. 2 mm long; disc 0.5–2 by 0.5–2 mm; stamens: anthers 2–2.7 by 0.2–0.4 mm long, terminal ap- pendage 0.5–1 mm long; pollen sacs 1.5–2 mm long; style 2.5–4.5 by 0.2–0.3 mm; stigma 2–3.5 by 0.5 mm. Fruit 3–6 by 4–7 mm, sparsely pubescent to pubescent (hairs 0.4–1 mm long, light brown to brown, or ginger-brown). Seeds 0.7–1 by 0.2–0.5 by 0.7 mm. Distribution — E Madagascar, Province Antsiranana, Re- gion Sava (Districts: Sambava, Andapa and Antalaha). Prov- ince Toamasina, Region Analanjirofo (Districts Maroantsetra, Mananara and Vavatenina); Region Alaotra-Mangoro (Districts Ambatondrazaka and Moramanga); Region Atsinanana (Dis- tricts Toamasina II and Brickaville). Province Fianarantsoa, Region Vatovavy-Fotovinany (District Ifanadiana). Habitat & Ecology —Based on specimen data: closed can­ opy, humid, evergreen forest; recorded on gneiss and clay; 10–850 m. Based on GIS data: humid forest; basement rocks, lavas and alluvial deposits; 100–1600 m; mean annual precipi- tation 1100–3000 mm; mean annual temperature 19–25 °C; dry season: usually 0–2 months but 7 months for population in Ambatondrazaka area. Map 1 Distribution of B. brevithyrsa A.P.Davis (-), B. crinita (A.Rich.)Wittle Phenology — Flowering: September to March; fruiting: No­ & A.P.Davis ($) and B. longithyrsa Baker (:). vember to May. 108 Blumea – Volume 55 / 2, 2010

Fig. 1 Bertiera crinita (A.Rich.) Wittle & A.P.Davis. a. Habit; b. part of shoot, showing interpetiolar stipule; c. flower subtended by bracteoles; d. cut and opened out corolla to show anthers, disc and style and stigma (pollen presenter); e. anther in adaxial view, showing apical connective appendage; f. immature fruit with persistent calyx lobes; g. transverse section of fruit, showing two locules, peltate placentas, and seeds; h. seed in two views (a, b: Rakotonasolo RNF 275; c–h: Davis et al. APD 1093; all K). Scale bars: a = 2 cm; b = 6 mm; c, d, f, g = 2 mm; e = 1 mm; h = 0.5 mm. P.M. Wittle & A.P. Davis: A revision of Madagascan Bertiera 109

Vernacular names — Sirontafoka (District Andapa); Tsirin­ Shrub to small tree, 1–5(–10) m high. Young shoots puberulous tafika (District Vavatenina); Soritafika and Hazovolena (District to pubescent (hairs 0.3–0.8 mm long, light brown to reddish Ambatondrazaka). brown). Stipules 6–20 by 3–5 mm, ± glabrous to puberulous Conservation status — IUCN Red List Category (IUCN, or sparsely pubescent, hairs mainly confined to central portion 2001): Near Threatened (NT). This species does not qualify of stipule (hairs 0.5–0.9 mm long, brown to reddish brown). for a threatened status (IUCN 2001) at present but is close to Leaves: petioles (1–)3–10 mm long, puberulous or pubescent qualifying in the future based on an area of occupancy (2 500 (like the young shoots); leaf blades elliptic to narrowly elliptic, km2 (based on a 10 × 10 km grid) and by having seven loca- or elliptic-ovate to elliptic-obovate, 9–22.5 by 2–7 cm; base tions (subpopulations). Other information: extent of occurrence narrowly cuneate to cuneate, sometimes rounded-cuneate, 53 844 km2. often slightly asymmetric; margin puberulous to very sparsely Notes — We originally proposed to describe a new species pubescent (hairs 0.2–0.3 mm long, light brown); apex attenuate for the above taxon but during a study of unplaced names in to attenuate-acuminate, rarely narrowly acute; lower surface: Rubiaceae (Ruhsam et al. 2008) we found that the unplaced midrib pubescent (hairs 0.4–0.6 mm long, appressed, light name Sabicea crinita A.Rich. represented our new species brown to brown); secondary veins 6–11 pairs, pubescent (like of Bertiera. We have deduced that the original material used the midrib but density and length of hairs less); tertiary venation by Richard (1830) is a sheet held in the Paris (P) herbarium. reticulate to ramified, obscure; higher order venation obscure; It bears the original label in the lower left-hand corner of the leaf surface (including secondary and tertiary venation) sparsely sheet, with “Sabicea crinita Nob.” written upon it. In another puberulous (hairs 0.2–0.3 mm long, brown); upper surface: hand, presumably written at a later date is written: “Herbarium midrib glabrous or very sparsely pubescent (hairs 0.4–0.7 Richard. Madagascar. Scripsit A.Richard.” On this sheet there mm long, light brown), venation and leaf surface glabrous. are clearly two elements: 1) a Bertiera sp., taking up most of Inflorescence an open, rather lax thyrse, 4.5–20 by 1.8–7(–8) the sheet, as detailed above; and 2) a much smaller specimen cm, all parts (peduncle, inflorescence branches, bracts and of a Mussaenda sp., circled in pencil and annotated “M. crinita bracteoles) puberulous to pubescent (hairs erecto-patent to A.M. H.” [A.-M. Homolle]. In the lower right-hand corner there appressed, 0.1–0.5(–0.7) mm long, light brown to brown, or is another label, which postdates the original. Printed on this reddish brown); peduncle 1–4.9 cm long; bracts present or label is “HERBIER E. DRAKE”, and written below (blue ink) in sometimes absent, if present narrowly triangular, linear, very an unknown hand is “Mussaenda crinita”. The original diagnosis narrowly elliptic, or very narrowly oblanceolate, 7–13(–22) by Richard (1830) clearly refers only to the Bertiera specimen: by 1–1.5(–2) mm; bracteoles narrowly triangular to narrowly “rufo-crinita; fol. elliptico-lanceolatis acuminatis praesertium lanceolate, 3–5.5 by 0.5–1 mm. Flowers: hypanthium 2.7–4.1 by 1.5–1.9 mm, puberulous to sparsely puberulous (hairs c. 0.2 subtus sparsè crinitis; stipulisque connatis acuminatis, acu- mm long, brown); calyx lobes triangular to narrowly triangular, mine longissimo; floribus terminalibus racemoso-congestis”, 0.3–1.5 by 0.3–0.5 mm; corolla 2.4–6 by 1.4–4 mm, exterior and not the smaller Mussaenda fragment. We have therefore sparsely puberulous, hairs mainly confined to median ridge of lectotypified Sabicea crinita based on the main element of the corolla lobes (hairs c. 0.1 mm long, light brown); corolla tube sheet and one that is unambiguously in accord with the diag- 2–3 by 1–2 mm; corolla lobes triangular, 1.5–2.3 by 1–1.5 nosis of Richard (1830). Homolle (1938) made the combination mm, apices acute to narrowly acute or very shortly acuminate, Mussaenda crinita (A.Rich.) Homolle, based on Sabicea crinita acumen 0.2–0.4 mm long; disc 0.5–1 by 0.5–1 mm; stamens: A.Rich., but instead of citing the original material (see above) anthers 1–1.5 by 0.3–0.5 mm, terminal appendage 0.3–0.5 a Chapelier specimen was cited (“Madagascar, Chapelier”; mm long; pollen sacs c. 0.8 mm long; style 2–2.5 by 0.2–0.3 Madagascar, Chapelier s.n. (holo P)). Richard (1830) cited mm; stigma c. 1.5 by 1 mm. Fruit 4–6 by 4–7 mm, glabrous or Chapelier specimens when they were used as original material, rarely very sparsely puberulous to pubescent (hairs 0.2–0.4 but in the case of Sabicea crinita he did not. mm long, light brown to brown). Seeds 0.7–1 by 0.2–0.7 by Bertiera crinita is very easily separated from B. longithyrsa 0.7–0.8 mm. and B. brevithyrsa, on the basis that it is a very much more Distribution — NW Madagascar, including the islands Nosy pubescent plant: the distribution, density and length of hairs Be and Nosy Komba, Province Antsiranana, Region Diana is considerably greater in B. crinita. Bertiera crinita also can (Districts Ambilobe, Ambanja and Nosy Be). Province Ma- be separated from B. longithyrsa due to its longer calyx lobes hajanga, Region Sofia (District Analalava). (1–4 mm vs 0.3–1.5 mm long), longer corolla tube (2.5–8 mm Habitat & Ecology — Based on specimen data: occurring vs 2–3 mm long) and generally shorter peduncle (0.2–3.5 cm in humid, evergreen forest (including Sambirano forest type) vs 1–4.9 cm long). Bertiera crinita also can be separated from and seasonally dry evergreen-deciduous forest, in primary B. brevithyrsa on the basis of its generally longer peduncle and secondary forest, frequently found adjacent to rivers and (0.2–3.5 cm vs 0.2–0.4 cm long), broader bracteoles (0.5–1.5 streams; one record on siliceous substrate (Humbert 18733 mm vs 0.2–0.4 mm wide) and generally smaller leaves (7.5– (P)); 0–1000(–1300) m. Based on GIS data: subhumid and 16.5 by 2–5.3 cm vs 4–10.4 by 1.1–2.9 cm). The distribution humid forest, woodlands; sandstones, basement rocks and of B. crinita and B. longithyrsa does not overlap (see Map 1). alluvial deposits; 100–1300 m; mean annual precipitation Their ecologies are also different, with the former in wetter for- 1400–2100 mm; mean annual temperature: 19–27 °C; dry est (higher mean annual precipitation, and shorter dry season; season: 6–7 months. see Habitat & Ecology, for each species). Bertiera crinita and Phenology — Flowering: (August–) September to Decem- B. brevithyrsa overlap in their distribution but do not appear to ber; fruiting: (August–) September to May. be sympatric. Vernacular names — Tsinitafiky (Districts Nosy Be, Anala- Bertiera crinita has been recorded as a climber (Herbier Institut lava); Duigaduigana (District Ambaja); Valomitiry (District Am­ Scientifique Madagascar 2489 (P); Cours 189[1] (P)) but this banja); Sevabe (District Ambanja). is erroneous. Conservation status — IUCN Red List Category (IUCN 2001): Vulnerable (VU). VU B1ab (i, ii, iii, iv, v); B1, extent of occurrence less than 20 000 km2 (B. longithyrsa: 6 757 km2); a, severely 3. Bertiera longithyrsa Baker — Map 1 fragmented or known to exist at 10 locations (the distribution Bertiera longithyrsa Baker (1890) 322. — Type: Baron 5788 (holo K), Mada- of B. longithyrsa is severely fragmented; known from 7 loca- gascar, Sambirano, Nossi-Bé [Nosy Be], 1887 (received at K). tions (subpopulations)); b (i–v), continuing decline inferred 110 Blumea – Volume 55 / 2, 2010 due to habitat loss and degradation. Other information: area of Nguembou CK, Sonké B, Zapfack L, Lejoly J. 2003. Les espèces camer- occurrence 2 100 km2 (based on a 10 × 10 km grid). Bertiera ounaises du genre Bertiera (Rubiaceae). Systematics and Geography of longithyrsa is confined to the NW region of Madagascar (Map 1). Plants 73: 237–280. Richard A. 1830. Mémoire sur la Famille des Rubiacées. Tastu, Paris. Most of the recent collections (post 1960) for this species have Robbrecht E, Manen JF. 2006. The major evolutionary lineages of the coffee been made in the southern areas of the range; the northern part family (Rubiaceae, angiosperms). Combined analysis (nDNA and cpDNA) has been reduced by agricultural development. For example, to infer the position of Coptosapelta and Luculia, and supertree construction suitable habitat near Ambilobe is either highly degraded or based on rbcL, rps16, trnL-trnF and atpB-rbcL data. A new classification reduced in extent; the last collection of B. longithyrsa from this in two subfamilies, Cinchonoideae and Rubioideae. Systematics and general locality was made in 1921. Geography of Plants 76: 85–146. Note — For details of the difference between B. longithyrsa Robbrecht E, Rohrhofer U, Puff C. 1994. A survey of Bertiera (Rubiaceae), including a discussion of its taxonomic position. Opera Botanica Belgica and B. crinita see Notes for the latter species (above). See 6: 101–141. Key to Species for differences between B. longithyrsa and Ruhsam M, Govaerts R, Davis AP. 2008. Nomenclatural changes in prepa- B. brevithyrsa. ration for a World Rubiaceae Checklist. Botanical Journal of the Linnean Society 157: 115–124. Steyermark JA. 1967. Bertiera. The botany of the Guayana Highland, VII. Acknowledgements This investigation would not have been possible Memoirs of the New York Botanical Garden 17, 1: 316–322. without the agreement of CAFF (Comité Ad’hoc Faune et Flore, which is the Schatz GE. 2001. Generic tree flora of Madagascar. Royal Botanic Gardens, association of the Ministère des Eaux et Forêts, Ministère de l’Enseignement Kew & Missouri Botanical Garden. Supérieure, Ministère de l’Environnement and Ministère de la Recherche Tosh J, Davis AP, Dessein S, De Block P, Huysmans S, Fay MF, Smets E, Scientifique, in Antananarivo, Madagascar). We would like to thank the Robbrecht E. 2009. Phylogeny of Tricalysia A.Rich. (Rubiaceae) and its directors and staff of the herbaria at K, MO, P, TAN and TEF for providing relationships with allied genera based on plastid DNA data: resurrection us with access to their collections. We also gratefully acknowledge the fol- of the genus Empogona. Annals of the Missouri Botanical Garden 96: lowing: Hazel Wilks for preparing the illustrations used in this contribution; 194–213. Franck Rakotonasolo for his assistance with herbarium specimens and Verdcourt B. 1983. Notes on Mascarene Rubiaceae. Kew Bulletin 37: 521– fieldwork; Mijoro Rakotoarinivo and Charlotte Cole for producing GIS data 576. for conservation assessments, map and ecological data; Sally Dawson for Wernham HF. 1912. A revision of the genus Bertiera. Journal of Botany 50: various support and advice. 110–117, 156–164. Willis F, Moat J, Paton A. 2003. Defining a role for herbarium data in Red REFERENCES List assessments: a case study of Plectranthus from eastern and southern tropical Africa. Biodiversity and Conservation 12: 1537–1552. Andreasen K, Bremer B. 2000. Combined phylogenetic analysis in the Rubiaceae-Ixoroideae: morphology, nuclear and chloroplast DNA data. IDENTIFICATION LIST American Journal of Botany 87: 1731–1748. Aublet JBC. 1775a. Histoire des Plantes de la Guiane François, vol. 1: 1 = Bertiera brevithyrsa 1–621. Didot Jeune, Paris. 2 = Bertiera crinita Aublet JBC. 1775b. Histoire des Plantes de la Guiane François, vol. 3: 3 = Bertiera longithyrsa 1–193. Didot Jeune, Paris. Baker JG. 1890. Further contributions to the Flora of Madagascar. Journal Anonymous (herb. Richard) s.n.: 2 — Anonymous 4878-RN: 3 — Antilahi- of the Linnean Society 25: 294–350. mena 46: 3 — Antilahimena et al. 448: 3. Bridson DM. 1988. Bertiera. In: Bridson DM, Verdcourt B, Rubiaceae, part 2, Baron 5788: 3 — Bernardi 11829: 3 — Birkinshaw & Andrews 28: 3 — Boivin Flora of Tropical East Africa: 479–483. Balkema, Rotterdam. 2070: 3 — Bosser 14742: 3 — Briggs et al. MB 125: 1. Bridson DM. 1998. Rubiaceae. In: Pope GV (ed), Flora Zambesiaca, vol. 5, Capuron 23705-SF: 2; 23921-SF: 2; 24937-SF: 2 — Capuron & Chauvet part 2: 211–377. Royal Botanic Gardens, Kew. 24776-SF: 2 — Cours 189 [1]: 2; 1891: 2; 4266: 2 — Cours & Humbert Bridson DM, Verdcourt B. 2003. Rubiaceae. In: Pope GV (ed), Flora Zambe- 5668: 3. siaca, vol. 5, part 3: 379–720. Royal Botanic Gardens, Kew. Davis & Rakotonasolo APD 4524: 1 — Davis et al. APD 1093: 2 — Déquaire Davis AP, Chester M, Maurin O, Fay MF. 2007. Searching for the relatives 27847: 2; 27975: 2 — Derleth 153: 3. of Coffea (Rubiaceae, Ixoroideae): the circumscription and phylogeny Gautier & Totozafy Be LG 2867: 3 — Gautier et al. LG 3187: 3; LG 3875: 2 of Coffeeae based on plastid sequence data and morphology. American — Gentry & Schatz 62214: 2. Journal of Botany 94: 313–329. Harizo 1264-RN: 3; 6904-RN: 3 — Hildebrandt 3002: 3 — Humbert 4006: 3; Hallé N. 1960. Sur les Bertiera (Rubiaceae) d’Afrique. Notulae Systematicae. 4015: 3; 18733: 3; 18767: 3; 21876 [a]: 2; 25929: 3 — Humbert & Capuron Phanerogramie (Paris) 16: 280–292. 24295: 2 — Humbert & Cours 32691: 3; 32907: 3. Hallé N. 1970. Bertiera. Rubiacées, 2e partie. In: Aubréville A, Leroy J-F Institut de Recherche Scientifique de Madagascar 2489: 2. (eds), Flore du Gabon vol. 17: 32–69. Muséum National d’Histoire Na- Jacquemin 453: 3 — Jardin Botanique de Tananarive 2760: 2; 2767: 2. turelle, Paris. Keraudren 1584: 3; 1600: 3; 1608: 3. Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A. 2005. Very high Laibosaka 7811-RN : 3 — Lowry et al. 4126b: 2. resolution interpolated climate surfaces for global land areas. International Malcomber 2791: 2; 2809: 2 — Malcomber et al. 1922: 3; 1969: 3; 2252: 3 Journal of Climatology 25: 1965–1978. — Morat 1408: 3. Holmgren PK, Holmgren NH, Barnett LC. 1990. Index Herbariorum. Part 1: Perrier de la Bâthie 3759: 3; 3833: 3; 3947: 2 — Pervillé 359: 3; 463: 3. The Herbaria of the World, 8th Edition. Regnum Vegetabile [series]. New Rabenantoandro et al. 457: 2 — Rabevohitra 4848: 2 — Raboarivo 7727-SF: York Botanical Garden, New York. 3 — Raharimalala 170: 2 — Rajokiny 11744-RN: 2 — Rakoto 4898-RN: Homolle A-M. 1938. Mussaenda noveaux de Madagascar. Notulae System- 3; 4899-RN: 3 — Rakotomalala & Velomaro 54: 3 — Rakotomalaza et al. aticae. Phanerogramie (Paris) 7: 3–7. 742: 2 — Rakotonasolo RNF 275: 2 — Rakotoniaina 2329-RN: 2; 2878- IUCN 2001. IUCN Red List Categories: Version 3.1. IUCN Species Survival RN: 2 — Rakotovao 10870-RN: 2 — Rakotozafy 275: 3 — Ramarokoto Commission, Cambridge. 7768-RN: 2 — Ranarivelo et al. RTI 292: 2 — Rasovimbahoaka 639: 2 Leroy J-F. 1974. Recherches sur la phylogenèse du développement. Mise — Razafindrazaka et al. 1: 3 — Reboroaka 7811-RN: 3 — Richard (herb. en évidence d’une série de trois états dans le genre Bertiera (Rubiacées). Richard) 173: 3 — Rokoalina [?] 7565-RN: 2. Adansonia, sér. 2, 14, 2: 53–59. Saboureau 58: 3 — Schatz 2749: 2 — Service Forestier de Madagascar Leroy J-F. 1989. Bertiera. In: Bosser J, Cadet T, Guého J, Marais W (eds), 3164-SF: 3; 3907-SF: 3; 26131-SF: 2. Flore des Mascareignes La Réunion, Maurice, Rodrigues. 107. Caprifolia- Tosh et al. JT 259: 2 — Totozafy Be STB 527: 3 — Tsilizy 6904-RN: 3. cées à 108bis. Valérianacées: 71–76. Mauritius Sugar Industry Research Waterlot 369: 3 — Wohlhauser & Ranjatson SW 60038: 3. Institute, Reduit, Mauritius; L’Office de la Research Scientifique et Tech- nique Outre-Mer, Paris, France; and Royal Botanic Gardens, Kew, UK.